Synergistic effect of Leclercia adecarboxylata CIP 82.92 and rice husk biochar on yield enhancement and cadmium reduction in mung bean

Synergistic effect of Leclercia adecarboxylata CIP 82.92 and rice husk biochar on yield enhancement and cadmium reduction in mung bean | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Synergistic effect of Leclercia adecarboxylata CIP 82.92 and rice husk biochar on yield enhancement and cadmium reduction in mung bean Nguyen Van Chuong, Tran Le Kim Tri, Nguyen Ngoc Phuong Trang, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7316646/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Mung bean ( Vigna radiata L.) is a vital legume crop widely grown in Asia, valued for its nutritional qualities and nitrogen-fixing ability. However, its productivity is challenged by cadmium (Cd) contamination in soils, which poses risks to crop yield and food safety. This study aimed to isolate indigenous endophytic bacteria capable of promoting mung bean growth while reducing Cd uptake, and to evaluate the combined effect of bacterial inoculation with rice husk biochar (RHB) in field trials conducted over two consecutive seasons (2023–2024 and 2024–2025) in Cd-contaminated soils of An Giang province, Vietnam. Leclercia adecarboxylata CIP 82.92 (strain CIP) was isolated from mung bean (MB) roots and characterized for morphological, biochemical, and molecular features, confirming its identity and nitrogen-fixing potential. Field experiments used a randomized complete block design with two factors: inoculation with CIP 82.92 and three RHB application rates (0, 5, and 10 t ha⁻¹), with chemical fertilizers uniformly applied. Results demonstrated that inoculation with CIP 82.92 significantly enhanced plant growth parameters, biomass, and yield components, while RHB application further improved these effects, showing a synergistic interaction. Importantly, Cd accumulation in seeds and aerial parts was markedly reduced by combined treatment, alongside improvements in seed protein and lipid content. The beneficial outcomes were more pronounced in the second year, indicating cumulative soil health benefits. This study highlights the integrated use of strain CIP 82.92 and RHB as an effective, eco-friendly approach to increase mung bean productivity and food safety in Cd-contaminated agricultural systems. Biochar Cadmium Indigenous bacteria Nitrogen-fixing Strain CIP 82.92 Figures Figure 1 Figure 2 Figure 3 Introduction Mung bean ( Vigna radiata L.) is one of the most important pulse crops in many Asian countries, including Vietnam, India, China, and Thailand, owing to its high nutritional value, short growth cycle, nitrogen-fixing ability, and adaptability to various agro-ecological conditions (Hou et al. 2019 ). As a valuable source of plant-based protein, vitamins, minerals, and bioactive compounds, mung bean plays a crucial role in ensuring food security, improving soil fertility, and supporting smallholder farmers' livelihoods in resource-limited regions (Kessy et al. 2024 ). In addition to its agronomic benefits, the crop contributes to sustainable agriculture by restoring soil nitrogen levels and reducing the need for synthetic fertilizers in crop rotation systems (Al-Shammary et al. 2024 ). However, in recent decades, the productivity and sustainability of mung bean cultivation have been increasingly threatened by a multitude of environmental stressors, most notably, climate change, soil degradation, and heavy metal contamination (Liu et al. 2025 ). The intensification of agricultural production has led to the overuse of chemical fertilizers and pesticides, contributing to declining soil health, reduced microbial diversity, and increased environmental pollution (Xing et al. 2025 ). Among these, cadmium (Cd) contamination has emerged as a particularly serious concern due to its high toxicity, persistence in soil, and ability to accumulate in edible plant parts (Chuong et al. 2024 ). Cd contamination in agricultural soils, originating from industrial discharge, phosphate fertilizers, and irrigation with polluted water, poses a dual threat: it not only reduces crop productivity but also endangers human health through dietary intake (Niño-Savala et al. 2019 ; Liang et al. 2024 ). MB, like many other legumes, is susceptible to Cd uptake and accumulation, which severely compromises seed quality and market value (Rehman et al. 2022 ; Davidova et al. 2024 ). Moreover, Cd stress negatively affects physiological and biochemical processes in plants, including nutrient uptake, photosynthesis, and enzymatic activity, ultimately leading to stunted growth and lower yields (Zhao et al. 2021 ; Rehman et al. 2022 ). This situation underscores the urgent need for environmentally friendly and effective approaches to mitigate Cd toxicity in mung bean cultivation, particularly in regions where soil contamination is widespread and traditional remediation practices are either too costly or impractical (Umer Chattha et al. 2021 ; Zulfiqar et al. 2023 ). In response to these challenges, the use of plant growth-promoting endophytic bacteria (PGPEB) in combination with organic soil amendments, such as biochar, has gained increasing attention as a sustainable strategy to enhance crop resilience, improve soil health, and reduce heavy metal uptake in plants (Kamyab et al. 2025 ; Chuong et al. 2025 ). Endophytic bacteria colonize plant tissues without causing disease and are known to confer multiple benefits to their host plants, including biological nitrogen fixation, phytohormone production, nutrient solubilization, and metal detoxification (Qadir et al. 2024 ). Among various PGPEB, nitrogen-fixing strains are particularly valuable in legumes, as they reduce dependence on synthetic nitrogen inputs and enhance plant growth under stressful conditions (Chuong 2024 a). RHB, a carbon-rich material produced by pyrolysis of biomass under limited oxygen conditions, has also been widely studied for its ability to immobilize heavy metals, improve soil structure, and support beneficial microbial communities. RHB is an abundant and low-cost by-product in many rice-producing countries (Asadi et al. 2021 ; Li et al. 2023 ). Its porous structure and high cation exchange capacity make it an effective soil amendment to reduce metal bioavailability and promote plant growth. When applied together, biochar and beneficial bacteria can create a synergistic effect, enhancing both microbial survival and plant performance under stress conditions (Kamyab et al. 2025 ). Despite these promising findings, relatively few studies have examined the combined effect of indigenous endophytic bacteria and biochar on mung bean productivity and Cd mitigation under field conditions. Furthermore, the potential of Leclercia adecarboxylata an emerging plant-associated bacterium with nitrogen-fixing potential remains largely unexplored in legume crops. Leclercia adecarboxylata has been identified in various plant rhizospheres and endophytic environments, showing promise in promoting growth and stress tolerance in host plants. However, its application in Cd-contaminated soils, particularly in mung bean cultivation, is novel and warrants detailed investigation (Meng et al. 2025 ) To address these knowledge gaps, the present study was undertaken to isolate and identify indigenous endophytic bacteria from mung bean roots and evaluate their potential in combination with RHB to enhance plant growth, yield, and nutritional quality while reducing Cd accumulation under field conditions. Specifically, we hypothesized that (1) inoculation with strain CIP 82.92 would improve mung bean performance through biological nitrogen fixation and growth stimulation; (2) application of rice husk biochar would reduce Cd bioavailability and enhance soil fertility; and (3) the combined use of biochar and bacterial inoculation would exert a synergistic effect, leading to significant improvements in yield and reduction of Cd accumulation in mung bean tissues. This study provides new insights into the integrated use of indigenous microbial resources and organic amendments to rehabilitate Cd-contaminated agricultural soils and promote sustainable mung bean production. The results contribute to a growing body of evidence supporting the ecological benefits of nature-based solutions in mitigating heavy metal stress and reducing dependency on synthetic agrochemicals. Materials and methods Collection and treatment of MB root samples The MB root samples were collected from MB cultivation fields in An Phu commune, An Giang province, Vietnam. This area is known to have Cd-contaminated soil, and previous studies have detected Cd accumulation in the stems, leaves, and seeds of the plants (Chuong et al. 2024 ). The roots were carefully uprooted, placed in sterile plastic bags, and transported to the laboratory. Prior to isolation, the MB roots were surface sterilized by thoroughly washing them with tap water to remove soil and debris. The MB roots were then immersed in 70% ethanol for 3 minutes for partial sterilization, followed by treatment with 2.5% sodium hypochlorite for 5 minutes to eliminate most surface bacteria. Finally, the roots were rinsed again with 70% ethanol and several times with sterile distilled water to remove any remaining sterilizing agents. To verify the effectiveness of surface sterilization, sterilized root segments were placed on agar plates. The absence of bacterial growth after incubation indicated successful sterilization (GohilK PatelH et al. 2020 ; Saldierna Guzmán et al. 2020 ). Isolation and Identification of Strain CIP 82.92 For the isolation of strain CIP 82.92, the surface-sterilized roots were homogenized and serially diluted up to 10⁻⁸. Aliquots (0.1 mL) from each dilution were spread onto yeast mannitol agar (YMA) plates and incubated at 32°C for 60 hours. Among the resulting colonies, ten isolates that exhibited morphological characteristics consistent with CIP 82.92 strain were selected for further study. These colonies were 3–4 mm in diameter, had smooth and regular margins, and were large, rod-shaped, endospore-forming, and Gram-positive as confirmed by Gram staining and microscopic observation at 100X magnification (Nxumalo et al. 2020 ). Molecular identification of the ten selected isolates was conducted using 16S rRNA gene sequencing. Amplification was performed using the universal primers 27F (5′-AGAGTTTGATCCTGGCTCAG-3′) and 1492R (5′-GGTTACCTTGTTACGACTT-3′) (Dos Santos et al. 2019 ; Hongmiao et al. 2021). The obtained sequences were compared with reference sequences in the GenBank database via BLAST. Phylogenetic analysis was carried out using MEGA 11 software, following the method of Saitou and Nei ( 1987 ) and Tamura et al. ( 2011 ), with 1,000 bootstrap replicates (Felsenstein, 1985). Ambiguous positions were excluded using the pairwise deletion option (Kumar et al. 2018 ). One colony was confirmed to be strain CIP 82.92, exhibiting 100% identity with reference strains. These sequences were subsequently submitted to the NCBI GenBank database and used to construct the phylogenetic tree (Fig. 1). Thermal adaptation Thermal adaptation of strain CIP 82.92 was conducted in two phases. Initially, four YMA nutrient medium test tubes were prepared for each strain. Subsequently, strain CIP 82.92 streaked onto YMA agar in the tubes and incubated at four different temperatures: 30, 35, 40, and 45°C. Each thermal level was tested in four replicates, and colony development was observed over the course of one week (Van Chuong and Le Kim Tri 2024 ). Salt adaptation The salt tolerance of the selected strain was evaluated using the following method: First, four test tubes containing YMA agar medium were prepared for each salt concentration. Then, NaCl was added to achieve final concentrations of 1.0%, 2%, 3%, 4%, and 5%. Selected colonies of the strain CIP 82.92 were subsequently inoculated into each tube and incubated at 28°C, with four replicates per concentration. Observations were recorded after one week (Mustafa A and Özden E 2023 ; Van Chuong and Le Kim Tri 2024 ). Ammonia production The ability of the identified strain CIP 82.92 to produce ammonia was evaluated qualitatively. Individual colonies were inoculated into peptone water and incubated on YMA medium at 30°C for 60 to 80 hours. Ammonia production was confirmed by a color shift from brown to yellow upon the addition of Nessler’s reagent (Borah et al. 2019 ). Nitrogenase activity Nitrogenase activity was determined using the acetylene reduction assay following the method described by Puri et al. ( 2018 ). The strain CIP 82.92 was initially cultured in YMA liquid medium and incubated for 24 hours. The resulting culture was then used to inoculate a nitrogen-free liquid medium. A control was maintained using the nitrogen-free medium without the addition of the strain CIP 82.92. The cell density of the CIP 82.92 suspension, measured spectrophotometrically at 600 nm, reached 0.8 under incubation conditions of 30°C and 160 rpm (Soper et al., 2021 ). Nitrogen concentration The strain CIP 82.92 was grown in a nitrogen-free medium supplemented with 0.05% malate, serving as the main carbon source, and incubated at a constant temperature of 30°C. After the incubation period, the cultures were centrifuged at 3000 rpm for one minute to separate the biomass. The resulting supernatant was carefully collected and used for nitrogen content analysis. This determination was conducted according to the procedure described by Daoliang et al. ( 2020 ), which allows for the accurate quantification of nitrogen fixed or released by the bacterial strains under nitrogen-limited conditions. Density augmentation of strain CIP 82.92 The selected strains were cultured in sterile YMA medium and incubated at 32°C for 60 hours. After incubation, the cultures were centrifuged at 6,000 rpm for 5 minutes, and the supernatant was removed. The bacterial cells were washed with sterile saline to adjust the concentration to 10⁸ CFU mL⁻¹. Suspensions of the five selected strains were then used to prepare a 1.0% (v/v) inoculum for evaluating plant growth-promoting factors. In the field experiment, 100 mL of this suspension was applied to MB seeds prior to sowing (Song et al. 2025 ). Preparation of MB variety The DX 208 MB variety has a growth duration of 70 to 80 days, with a determinate growth habit. The average yield depends on the level of cultivation intensity. The seeds have a green, moldy appearance that meets consumer preferences. DX 208 exhibits synchronous maturity, making it convenient for harvesting, which is typically done 2 to 3 times per cropping season. The variety shows good lodging resistance and moderate to high resistance to Cercospora leaf spot and powdery mildew. DX 208 is well-suited for cultivation in delta and coastal regions within crop rotation systems ( https://sonnptnt.ninhthuan.gov.vn ) Field Eexperimental design. NPK fertilization for the 2023–2024 and 2024–2025 seasons: Fertilizers were applied at rates of 40 kg N, 60 kg P₂O₅, and 50 kg K₂O per hectare, divided into three stages as follows: (i) Basal application: The entire amount of P₂O₅, along with one-third of the N and one-third of K₂O, was applied before sowing; (ii) First topdressing (at the 3-true-leaf stage): An additional one-third of N and one-third of K₂O was applied; (iii) Second topdressing (25–30 DAS): The remaining amounts of N and K₂O were applied (Trang and Chuong, 2024 ). Field experiments were conducted in An Phu commune, An Giang province, Vietnam, located at approximately 10.845°N latitude and 105.080°E longitude, from October to January of the 2023–2024 and 2024–2025 seasons. The experimental site’s climate transitions from the rainy season to the dry season during this period. From October to November per year, the area experiences high rainfall and humidity, whereas December to January is dry with little rainfall and temperatures ranging from 22–30°C, characterized by cool and stable weather. A randomized complete block design with two factors was used for both crop years (October to January of 2023–2024 and 2024–2025): Factor (1) with and without inoculation of strain CIP 82.92, and Factor (2) three levels of RHB application: 0.0, 5.0, and 10.0 t ha⁻¹. This resulted in six treatment combinations, each replicated four times. Inorganic fertilizers (40 kg N, 60 kg P₂O₅, and 50 kg K₂O per hectare) were uniformly applied to all treatments (Table 1 ). Table 1 Treatments of strain CIP 82.92 inoculation and different RHB rates in 2023–2024 and 2024–2025 Treatments Strain CIP 82.92 (10 8 CFU mL − 1 ) RHB (t ha − 1 ) Chemical fertilizer (kg ha − 1 ) Years 2023–2024 2024–2025 2023–2024 2024–2025 MB1 No No 0 0 40N- 60P 2 O 5 -50 K 2 O MB2 No No 5 5 MB3 No No 10 10 MB4 Yes Yes 0 0 MB5 Yes Yes 5 5 MB6 Yes Yes 10 10 Three pre-germinated mung bean seeds were sown per hole. Irrigation was carried out using water sourced from the Mekong River Delta. The experiment consisted of 6 treatments (Table 1 ), each replicated four times, with a total experimental area of 480 m² (2 m width × 10 m length × 4 replications × 6 treatments). MB seeds were sown at a spacing of 25 cm × 30 cm, with two seeds per hole, and thinned to retain one healthy plant 15 days after sowing (DAS). All treatments received chemical fertilizers at rates of 40 kg urea ha⁻¹, 60 kg P₂O₅ ha⁻¹, and 50 kg K₂O ha⁻¹, supplied by Binh Dien Fertilizer Company (Vietnam). Twenty-four soil samples were collected from the experimental treatments 15 days before the start of the experiment and analyzed, with the results presented in Table 2 . Agronomic parameters such as plant height, number of shoots, and chlorophyll content were monitored 65 days after sowing. Yield components, total yield, lipid and protein contents, as well as Cd accumulation in the stems, leaves, and seeds of MB were assessed annually at harvest. Cadmium concentrations were determined using atomic absorption spectrophotometry. Plant and soil analyses were conducted at the laboratories of An Giang University and Eurofins Company in Can Tho Province, Vietnam. Table 2 Physicochemical properties of farmland soil and RHB prior to the experiment (n = 24) Property Result Property Result Sand (%) 60.0 Total N (%) 0.102 Silt (%) 36.0 Available P (mg kg − 1 ) 72.7 Clay (%) 4.00 Exchangeable K (mg kg − 1 ) 106 pH soil 6.00 SOM (%) 2.21 Cadmium in soil (µg kg − 1 ) 123 CEC (cmol kg − 1 ) 6.02 2.11 Data collection and analysis Agronomic and yield parameters in both years, including total biomass, number of filled pods, filled pod weight, 1000-seed weight, were recorded at harvest. Fresh pod yield (t ha⁻¹) was determined for each treatment in both experimental years. In addition, seed samples were subjected to biochemical analyses to quantify lipid, protein, and Cd concentration of MB stems, leaves and seeds, following standard analytical protocols. All collected data were statistically analyzed using Statgraphics XV and Microsoft Excel. Analysis of variance (ANOVA) was performed, and treatment means were compared using the Least Significant Difference (LSD) test at a 5% significance level (p ≤ 0.05). Results Biochemical and genetic profiling of strain CIP 82.92 The colony of strain CIP 82.92 was selected from 20 initially isolated pure colonies based on Gram staining results and morphological characteristics, followed by molecular identification. The selected strain, Leclercia adecarboxylata CIP 82.92, was confirmed to share 100% genetic similarity with Leclercia adecarboxylata NBRC 102595 (Fig. 2 ). In the next step, its morphology was further examined to assess its nitrogen-fixing potential prior to conducting field experiments. The VITEK 2 system (BioMérieux, France) provided a 92% probability of identification and was further confirmed by Illumina-based genetic analysis. This system was employed to characterize the morphological and biochemical features of the four selected strains. Morphologically, prior to molecular identification, strain CIP 82.92 appeared as creamy white colonies with a diameter of approximately 3 mm, lacking pigmentation, and exhibiting shapes ranging from circular to irregular with smooth to undulate margins on YMA medium (Fig. 1). As shown in Fig. 1 shows that strain CIP 82.92 was identified as a Gram-positive, rod-shaped, motile bacterium, catalase-positive, and capable of aerobic growth at temperatures ranging from 20 to 50°C. Optimal growth occurred at pH 4–8, within the same temperature range, and in media containing up to 5% NaCl. Strain B was primarily identified based on phenotypic characterization and partial results obtained from the VITEK 2 system, which are summarized in Table 3 . Furthermore, phylogenetic analysis based on 16S rRNA gene sequences, with a total branch length of 0.001, confirmed 100% genetic similarity with the reference sequence of Leclercia adecarboxylata NBRC 102595, as illustrated in Fig. 2 . Table 3 The physical and chemical characterization of strains CIP 82.92 was determined using the VITEK 2 analyzer Biochemical test strains CIP 82.92 Biochemical test strains CIP 82.92 Beta–Xyloidine + NaCl (1–5%) ++ D–Mannose - Temperature (20-50 0 C) + D–Glucose - pH (4.0–8.0) ++ D–Galactose - Citrate use + D–Ribose - β – Glucosidase + Catalase ++ Mannitol + Oxidase - Raffinose + Starch hydrolysis + Glucose oxidation + (-): negative reaction; (+) weak reaction; (++) strong reaction 3.2 Assessment of ammonia activity and concentration The acetylene reduction to ethylene (C₂H₄) process, monitored via the C₂H₂ gas injection system, initially exhibited a short lag phase after acetylene introduction and gradually stabilized over the 72-hour incubation period. As shown in Fig. 3 , both nitrogenase activity and nitrogen concentration increased progressively with inoculation time. Specifically, the nitrogenase activity began at a low level (approximately 20 nmol C₂H₄/h/mL at 8 hours) and steadily rose, peaking at around 380 nmol nmol C₂H₄ h − 1 mL − 1 after 72 hours. Similarly, nitrogen concentration measured by the Kjeldahl method followed a comparable upward trend, increasing from below 50 mg L − 1 at 8 hours to over 420 mg L − 1 at 72 hours. Findings confirm that the selected strain CIP 82.92 actively participates in biological nitrogen fixation, with clear evidence of sustained nitrogenase enzyme activity over time. The high correlation between ethylene production and nitrogen accumulation suggests that this strain maintains robust diazotrophic capacity under laboratory conditions. Effect of strain CIP 82.92 and RHB on MB agronomic traits . The data presented in Table 4 clearly demonstrates the positive influence of both strain CIP 82.92 and RHB on the agronomic traits of mung bean, including plant height, number of branches per plant, and chlorophyll index, measured at 65 DAS. Across both years, inoculation with strain CIP 82.92 significantly improved all measured parameters. In the 2024–2025 season, inoculated plants reached a mean height of 67.5 cm compared to 60.5 cm in non-inoculated controls, while branch number increased from 12.0 to 17.9, and chlorophyll index rose dramatically from 43.9 to 57.6. Application of RHB at increasing rates also significantly improved mung bean growth. The highest application rate (10.0 t ha⁻¹) resulted in the greatest increases in all parameters in both years, especially in 2024–2025 where plant height and chlorophyll index peaked at 74.5 cm and 59.6, respectively. Significant interaction effects [F(A×B)] were observed, indicating a synergistic relationship between CIP 82.92 and RHB. For example, combined treatment consistently resulted in greater improvements than either factor alone. Notably, the improvements were more pronounced in the 2024–2025 season across all treatments. Biomass accumulation is a direct indicator of plant vigor and photosynthetic efficiency. According to the results, both the application of Bacillus strain CIP 82.92 and rice husk biochar (RHB) significantly increased the biomass of mung bean in both cropping years, with a notably stronger response in the 2024–2025 season. Inoculation with CIP 82.92 increased biomass from 20.2 g plant⁻¹ to 25.5 g plant⁻¹ in 2023–2024, and from 20.1 g to 30.5 g plant⁻¹ in 2024–2025. The more substantial increase in the second year (a 52% rise versus 26% in the first year) suggests that environmental conditions or cumulative soil microbiome enhancement may have amplified the bacterial effect. RHB at 10.0 t ha⁻¹ led to the highest biomass in both seasons, increasing from 28.7 g (2023–2024) to 40.8 g plant⁻¹ (2024–2025). Even at 5.0 t ha⁻¹, RHB significantly enhanced plant biomass over the control. The combined effect of CIP 82.92 and RHB was statistically significant [F(A×B)] at p ≤ 0.01, indicating a synergistic interaction. Notably, the biomass response was consistently greater in 2024–2025 across all treatments, suggesting long-term soil conditioning or climatic factors improved microbial establishment and root-soil interactions. Table 4 Effect of strains CIP 82.92 and RHB on MB-agronomic components at 65 DAS Factors Plant height (cm) Branch number (branches plant − 1 ) Chlorophyll index Biomass (g plant − 1 ) Years 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 Strains CIP 82.92 (A) No 61.5b 60.5b 12.3b 12.0b 44.9b 43.9b 20.2b 20.1b Yes 64.5a 67.5a 15.9a 17.9a 47.6a 57.6a 25.5a 30.5a RHB application (B) 0.0 t ha − 1 61.3c 61.3b 12.75b 12.7b 44.0b 42.0b 20.1b 19.6c 5.0 t ha − 1 63.1b 73.1ab 15.6a 17.6a 48.3a 58.3a 28.5a 33.5b 10.0 t ha − 1 64.5a 74.5a 15.5a 17.5a 49.6a 59.6a 28.7a 40.8a F (A) ** ** ** ** ** ** ** ** F (B) ** ** ** ** ** ** ** ** F (A x B) ** ** ** ** ** ** ** ** Note: ± indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at the 1% levels. Effect of strain CIP 82.92 and RHB on MB yield traits and productivity Table 5 Effect of strains CIP 82.92 and RHB on MB yield trait and productivity Factors Pod number (Pods per plant) Filled pod weight (g plant − 1 ) Weight of 1, 000 seeds (g) Fresh yield (t ha − 1 ) Years 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 Strains CIP 82.92 (A) No 35.7b 30.1b 18.5b 16.5b 35.2b 32.2b 1.5c 1.61c Yes 45.7a 54.1a 25.8a 27.8a 39.1a 41.1a 2.1a 2.42a RHB application (B) 0.0 t ha − 1 33.4c 32.1c 16.8c 17.8b 33.2c 33.2b 1.3c 1.9c 5.0 t ha − 1 43.2b 50.2b 24.7b 28.7a 45.2b 49.2a 2.0a 2.21b 10.0 t ha − 1 45.7a 55.6a 27.0a 28.0a 47.4a 49.4a 1.97a 2.54a F (A) ** ** ** ** ** ** ** ** F (B) ** ** ** ** ** ** ** ** F (A x B) ns ** ns ** ns ** ** ** Note: ± indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at P > 0.01. Table 5 proves that application of CIP 82.92 significantly increased pod numbers from 35.7 to 45.7 pods/plant in 2023–2024 and even more so in 2024–2025, from 30.1 to 54.1 pods/plant, reflecting a 79.7% increase in the second year. Similarly, RHB at 10.0 t ha⁻¹ resulted in the highest pod numbers (45.7 and 55.6 pods plant − 1 in 2023–2024 and 2024–2025, respectively). Significant enhancements were recorded with both treatments. Strain CIP 82.92 increased filled pod weight by 39% (2023–2024) and 68% (2024–2025). RHB also showed consistent increases, especially at 5.0–10.0 t ha⁻¹ (up to 28.7 g plant − 1 ). The MB pod number and weight are critical yield components and market determinants. CIP 82.92 increased 1000-seed weight by about 11% (2023–2024) and 28% (2024–2025). Fresh yield is the ultimate productivity measure. Inoculation with CIP 82.92 raised yield from 1.5 to 2.1 t ha⁻¹ in the first year and from 1.61 to 2.42 t ha⁻¹ in the second. RHB significantly enhanced yield, especially at 10.0 t ha⁻¹, reaching 2.54 t ha⁻¹, the highest recorded. Impact of strain CIP 82.92 and RHB nutrient compositions and Cd accumulation in MB seeds Both strain CIP 82.92 and RHB significantly increased the lipid and protein contents of MB seeds over the two years. Notably, CIP 82.92 increased lipid content from 25.2–26.8% in 2023–2024 and 25.3–27.8% in 2024–2025, while protein content improved from 17.3–18.0% and 17.1–19.0%, respectively. RHB application showed a dose-dependent improvement, especially at 10 t ha⁻¹, with lipid content reaching 31.9% and 32.3%, and protein content 18.2% and 19.2% in the two years, respectively (Table 6 ). Cd accumulation in both plant parts was significantly reduced by treatments with CIP 82.92 inoculation and RHB application. inoculation of strain CIP 82.92 reduced Cd in seeds from 61.7 to 36.1 µg kg⁻¹ (2023–2024) and from 77.6 to 23.4 µg kg⁻¹ (2024–2025). A 62–70% reduction, confirming its Cd-immobilizing and phytoprotective properties. The Cd concentrations in stems and leaves also dropped significantly under CIP 82.92, indicating systemic mitigation. RHB application at 5.0 t ha⁻¹ reduced seed Cd to 23.4 and 11.9 µg kg⁻¹, while 10.0 t ha⁻¹ lowered it to 37.6 µg kg⁻¹ in 2023–2024 and undetectable levels in 2024–2025. Importantly, the interaction effect [F(A×B)] was significant for Cd reduction, particularly in seeds, indicating that co-application of Strains CIP 82.92 and RHB is more effective than either alone. The co-application of CIP 82.92 and RHB significantly enhanced the nutritional quality of mung bean seeds while drastically reducing cadmium accumulation, particularly in the second year. These findings support the integration of endophytic bacteria and biochar as a sustainable, eco-friendly strategy for safe legume production on Cd-contaminated soils (Table 6 ). Table 6 Effect of strain CIP 82.92 and RHB on nutrient traits and Cd accumulation in MB seeds Factors Lipid Protein Cd concentration (µg kg − 1 ) Seeds (%) Stems and leaves Seeds Years 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 2023–2024 2024–2025 Strains CIP 82.92 (A) No 25.2 b 25.3b 17.3 b 17.1 b 90.9 a 108a 61.7 a 77.6 a Yes 26.8a 27.8a 18.0 a 19.0 a 62.2 b 53.0b 36.1b 23.4 b RHB application (B) 0.0 t ha − 1 28.4c 27.4b 16.9b 17.3b 159 a 117 a 77.6 a 87.6a 5.0 t ha − 1 31.1b 32.1a 17.9a 19.9a 65.2 b 63.0b 23.4 b 11.9b 10.0 t ha − 1 31.9a 32.3a 18.2a 19.2a 64.7 b 59.1 c 37.6 c undetected F (A) ** ** ** ** ** ** ** ** F (B) ** ** ** ** ** ** ** ** F (A x B) ** ns ns ** ns ** ** ** Note: ± indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at P < 0.01. Discussion Biochemical and genetic profiling of strain CIP 82.92 The strain CIP 82.92, a Gram-positive, rod-shaped bacterium, was isolated from MB roots cultivated in cadmium-contaminated soils using YMA medium. Biochemical assays confirmed positive catalase and oxidase activities, consistent with the typical characteristics of endophytic nitrogen-fixing bacteria reported in recent studies (Hossain et al. 2023 ). The strain demonstrated moderate salt tolerance, thriving in media containing 1–5% NaCl, which aligns with findings that most endophytes adapt well to low-to-moderate salinity conditions (Chandran et al. 2021 ; Khushboo et al. 2023; Falfán-Cortés et al. 2022 ). Molecular identification based on 16S rRNA gene sequencing revealed 100% similarity with known nitrogen-fixing bacteria, confirming the taxonomic position of the strain as Leclercia adecarboxylata , a species recognized for its growth-promoting capacity and cadmium resistance (Wang et al. 2024 ). The optimal growth temperature ranged from 20°C to 50°C, with 37°C being ideal for symbiotic nitrogen fixation, consistent with earlier reports on endophyte thermotolerance (Van Chuong and Le Kim Tri 2024 ; Nguyen Van 2025 ). Effect of strain CIP 82.92 and RHB on MB agronomic traits Recent studies have emphasized that Leclercia adecarboxylata not only plays a crucial role in nitrogen fixation under abiotic stress conditions but also promotes plant development via phytohormone production, phosphate solubilization, and heavy metal resistance demonstrating its potential as a biofertilizer in sustainable agriculture (Woo et al. 2025 ). These results are in accordance with previous findings confirming the effectiveness of the acetylene reduction assay (ARA) as a reliable indicator of nitrogenase activity and biological nitrogen fixation capacity (Aasfar et al. 2024 ). This supports prior findings that biochar enhances soil structure, cation exchange capacity, and microbial habitat (Torita et al., 2022 ). This synergy suggests that biochar not only improves the physical-chemical properties of soil but also serves as a carrier or habitat for beneficial microbes, enhancing their survival and colonization efficiency. Similar interactions have been reported by Cao et al. ( 2021 ), who observed that biochar-amended soils facilitated root colonization by endophytic bacteria, leading to improved nutrient uptake and growth. This aligns with the nitrogen-fixing potential of CIP 82.92, as previously confirmed through acetylene reduction assays (Fig. 3 ), and suggests more efficient nitrogen use efficiency when both amendments are applied in combination (Barbosa et al., 2023 ). The combined application of strain CIP 82.92 and RHB significantly enhanced MB growth attributes in both years, with stronger responses observed in the second season. These results highlight the potential of integrating PGPR and biochar as a sustainable and synergistic strategy for legume crop improvement. The high nitrogenase activity of strain CIP 82.92, as indicated by the ARA, reflects its strong potential for biological nitrogen fixation. This observation is consistent with recent reports on Leclercia adecarboxylata being isolated from leguminous crops (Meng et al. 2023 ; Woo et al. 2025 ). The elevated nitrogen levels recorded in the essay further confirm the strain’s effectiveness in enhancing nitrogen bioavailability, is a key factor in crop productivity. This effect likely stems from the strain's ability to adapt to the rhizosphere environment and to establish symbiotic interactions with legume roots (Concha et al. 2020; Chuong et al. 2024 ). Its nitrogen-fixing efficiency is comparable to, or even surpasses, other known endophytic diazotrophs, underscoring its potential as a biofertilizer candidate (Figiel et al. 2025 ). Effect of strain CIP 82.92 and RHB on MB yield trait and yield The application of strain CIP 82.92 in combination with RHB significantly improved nutrient availability and vegetative growth in MB. This is supported by recent studies highlighting RHB’s role in enhancing soil structure and nutrient uptake in legumes (Ben Gaied et al. 2024 ). The observed plant growth-promoting effects of CIP 82.92 may be attributed to mechanisms such as phytohormone production, phosphate solubilization, and nitrogen fixation, as also reported by Etesami ( 2025 ) and Flores et al. ( 2025 ). The interactive effect between the microbial inoculant and organic amendment underscores the synergistic benefits of integrating beneficial microbes with organic inputs, a strategy widely advocated in integrated nutrient management systems (Folina et al. 2025 ; Liang et al. 2025 ). Field trials conducted over two consecutive years revealed that combined application of CIP 82.92 and RHB at 10 t ha⁻¹ significantly enhanced MB yield and quality parameters compared to uninoculated or singly treated controls. Filled pod number reached 176 pods per plant and filled pod weight increased to 196 g per plant, representing 31% and 65% increases, respectively, over untreated controls. In contrast, the number and weight of unfilled pods decreased, indicating more efficient reproductive development and resource partitioning (LSD ≤ 0.01). These results are consistent with prior studies highlighting the role of endophytic bacteria and biochar in improving flower development, nutrient acquisition, and carbon allocation to reproductive organs (Villa-Parejo et al., 2025 ). RHB applications were also shown to improve soil physical properties and nutrient retention, supporting better seed development (Trozzo et al., 2025 ). Thousand-seed weight increased to 47.4–49.4 g under the 10t RHB ha⁻¹ treatment, reflecting improved assimilate allocation for seed filling (Zhu et al., 2025 ). These findings are consistent with recent reports demonstrating that RBH enhances nutrient availability and soil microbial activity, leading to increased growth and yield of leguminous crops (Abdul-Aziz et al. 2025 ). Biochar is known to enhance aeration, water retention, and microbial habitat quality in soil, particularly after multi-season application (Kumar et al., 2021 ). Recent studies further indicate that RHB combined with endophytic nitrogen-fixing bacteria significantly improves nodule formation and legume productivity, including in mung bean compared to treatments with low fertilizer or no inoculants (Linh et al. 2025 ). Notably, microbial inoculation often exerts greater influence on yield improvement than nitrogen fertilizers alone, emphasizing the value of biological inputs in sustainable legume production systems (López-Román et al.2025). In addition to boosting yield, the RHB-bacteria combination enhances soil health by increasing organic matter and beneficial microbial populations, which support long-term soil fertility (Ali et al. 2025 ). The integration of CIP 82.92 with RHB offers a promising strategy for increasing mung bean yield and seed protein content while concurrently reducing cadmium accumulation in edible plant parts. This approach contributes to sustainable agriculture through improved soil fertility, reduced environmental impact of synthetic fertilizers, and potential biocontrol of root diseases such as Phytophthora root rot via microbial antagonism and enhanced plant health (Woo et al. 2025 ). The use of biofertilizer and RHB combinations is recommended for MB cultivation on nutrient-depleted and Cd-contaminated soils to maximize productivity, farmer income, and environmental sustainability. The observed effects may result from the strain’s nitrogen-fixing and phosphate-solubilizing capabilities, which facilitate more efficient nutrient uptake consistent with recent findings on endophytic bacteria in sustainable farming systems (Dadhich et al. 2025 ). Effect of strain CIP 82.92 and RHB on MB nutrition and Cd accumulation These results indicate that CIP 82.92 promotes plant growth, possibly through enhanced nitrogen fixation and phytohormone synthesis consistent with previous studies on beneficial endophytes such as Bacillus aryabhattai and Bacillus subtilis (Radhakrishnan et al., 2017 ; Chuong et al., 2024 b). Biochar’s known capacity to improve soil aeration, moisture retention, and nutrient availability also provides favorable conditions for microbial colonization (Chen et al., 2018 ). The synergistic effect of biochar and PGPR has been shown to increase photosynthetic efficiency and biomass accumulation in legumes (Zhang et al., 2024 ). Stronger plant responses observed in the second year (2024–2025) may reflect cumulative benefits from biochar conditioning, microbial establishment, and improved soil health. These findings are supported by studies reporting that repeated applications of biochar and PGPR enhance rhizosphere stability and functional resilience (Zhang et al., 2024 ; Trang and Chuong, 2024 ). The greater reduction in Cd uptake and higher nutrient concentrations observed in 2024–2025 may be due to long-term improvements in soil structure, microbial colonization, and Cd immobilization. This supports biochar’s role in enhancing nutrient availability and microbe–root interactions, as previously reported (Ahmad et al. 2024 ). Potential mechanisms include extracellular polymeric substance (EPS) production, biosorption, and modified cadmium translocation pathways—consistent with recent studies on endophytic bacteria (Haider et al. 2022 ). Collectively, these findings confirm that biochar functions as a strong sorbent, binding heavy metals in soil and reducing plant uptake (Ma et al. 2024 ). Conclusion Based on the conducted field experiments over two consecutive seasons (2023–2024 and 2024–2025) in Cd-contaminated soils of An Phu commune. The synergistic application of strain CIP 82.92 and RHB significantly enhanced MB growth, yield, and seed quality while reducing Cd accumulation. The inoculation with strain CIP 82.92 improved plant height, branch number, chlorophyll content, biomass accumulation, pod numbers, filled pod weight, and 1,000-seed weight compared to uninoculated controls. Increasing RHB rates (up to 10 t ha⁻¹) also positively affected all growth and yield parameters, with maximal improvements observed under combined treatments, indicating a strong synergistic interaction. Importantly, the combined treatment substantially reduced cadmium concentrations in seeds and aerial plant parts, enhancing food safety. Seed lipid and protein contents were elevated with both CIP 82.92 and RHB treatments, further improving seed nutritional value. The beneficial effects are attributed to enhanced nitrogen fixation, phytohormone production from CIP 82.92, and improved soil physicochemical properties and microbial habitats provided by RHB. The more pronounced benefits observed during the second year suggest cumulative improvements in soil health and microbial establishment. This study demonstrates the potential of integrating endophytic bacteria and biochar amendments as an effective agronomic strategy to increase legume productivity and mitigate heavy metal stress in contaminated soils. Declarations Author contributions: Tran Thanh Liem, Phan Tran Hai Dang, Tran Le Kim Tri and Nguyen Ngoc Phuong Trang conducted experiments, collected data, performed chemical analyses, and assisted with statistical analysis. Nguyen Van Chuong conceived and designed the study, supervised the project, analyzed data, drafted and finalized the manuscript for submission. Funding No funding was received to assist with the preparation of this manuscript Data availability No datasets were generated or analyzed during the current study Code availability Not applicable Declarations Ethical approval Not applicable. 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Int J Mol Sci 25:13702. https://doi.org/10.3390/ijms252413702 Woo J-I, Adhikari A, Gam H-J, Jeon JR, Lee D-S, Kwon E-H, Kang S-M, Yun B-W, Lee I-J (2025) Integrated role of biochar and PGPR ( Leclercia adecarboxylata HW04) in enhancing cadmium phytoremediation and stress tolerance in Glycine max L. Plant Physiol Biochem 220:109489. https://doi.org/10.1016/j.plaphy.2025.109489 Woo JI, Adhikari A, Gam HJ, Jeon JR, Lee DS, Kwon EH, Kang SM, Yun BW, Lee IJ (2025) Integrated role of biochar and PGPR ( Leclercia adecarboxylata HW04) in enhancing cadmium phytoremediation and stress tolerance in Glycine max L. Plant Physiol Biochem 220:109489. https://doi:10.1016/j.plaphy.2025.109489 Xing Y, Xie Y, Wang X (2025) Enhancing soil health through balanced fertilization: a pathway to sustainable agriculture and food security. Front Microbiol 16:1536524. https://doi:10.3389/fmicb.2025.1536524 Zhang W, Niu W, Luo H (2024) Effect of biochar amendment on the growth and photosynthetic traits of plants under drought stress: a meta-analysis. Agronomy 14(12):2952. https://doi.org/10.3390/agronomy14122952 Zhao H, Guan J, Liang Q, Zhang X, Hu H, Zhang J (2021) Effects of cadmium stress on growth and physiological characteristics of sassafras seedlings. Sci Rep 11:9913. https://doi.org/10.1038/s41598-021-89322-0 Zhu S, Guo Y, Zhou, Luo W, Yi X, Zhou Y, Wu Y, Daniel F, Petticord DF, Song X (2025) Biochar efficacy in enhancing soil carbon fractions is mediated by parent soil type in grazing karst grassland. Carbon Res 4:1. https://doi:10.1007/s44246-025-00222-8 Zulfiqar U, Maqsood MF, Mohy-Ud-Din W, Shabaan M, Ahmad M, Kaleem M, Ishfaq M, Aslam Z, Shahzad B (2023) Recent advances in microbial-assisted remediation of cadmium-contaminated soil. Plants 12:3147. https://doi:10.3390/plants12173147 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7316646","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":497519732,"identity":"743eac05-a5cc-4e30-a45c-755c4996f887","order_by":0,"name":"Nguyen Van Chuong","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAzklEQVRIiWNgGAWjYHACNjDJD+Ewk6BFsoFkLQYHiNViLpH87DFPxR27zcePX5NgqLBObJDIPYBXi+WMNHNjnjPPkredySmTYDiTDtSSl4BXi8GZM2zSvG2Hk80O5KRJMLYdTmzgOWNAnBbj/jdALf+I0XK8B6zFzkAi/ZgEYwNQC3sPIS1tZpJzzhxOkLjxhtki4Vi6cRtBLYeZn0m8qThsz9+f/vDGhxpr2X5mHvxaYCCxgQGoMoEBFk1EAHsGBvYHxCoeBaNgFIyCEQYAoVdEsu6ZVYcAAAAASUVORK5CYII=","orcid":"","institution":"An Giang University","correspondingAuthor":true,"prefix":"","firstName":"Nguyen","middleName":"Van","lastName":"Chuong","suffix":""},{"id":497519733,"identity":"5cbf4aa0-f92c-47cd-94f1-84e45ef5dfa7","order_by":1,"name":"Tran Le Kim Tri","email":"","orcid":"","institution":"An Giang University","correspondingAuthor":false,"prefix":"","firstName":"Tran","middleName":"Le Kim","lastName":"Tri","suffix":""},{"id":497519737,"identity":"47932de9-865e-43f1-8fa4-340e7d2cd94c","order_by":2,"name":"Nguyen Ngoc Phuong Trang","email":"","orcid":"","institution":"An Giang University","correspondingAuthor":false,"prefix":"","firstName":"Nguyen","middleName":"Ngoc Phuong","lastName":"Trang","suffix":""},{"id":497519739,"identity":"53a9f62f-1ecd-4565-9b7b-f8cf03771870","order_by":3,"name":"Tran Thanh Liem","email":"","orcid":"","institution":"An Giang University","correspondingAuthor":false,"prefix":"","firstName":"Tran","middleName":"Thanh","lastName":"Liem","suffix":""},{"id":497519740,"identity":"8f943c7e-4c31-4126-9fb2-efe6343db136","order_by":4,"name":"Phan Tran Hai Dang","email":"","orcid":"","institution":"An Giang University","correspondingAuthor":false,"prefix":"","firstName":"Phan","middleName":"Tran Hai","lastName":"Dang","suffix":""}],"badges":[],"createdAt":"2025-08-07 08:53:15","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7316646/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7316646/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":88812351,"identity":"80b5f63b-6fde-47e2-bf7e-f447b6d56c68","added_by":"auto","created_at":"2025-08-11 15:43:45","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":439996,"visible":true,"origin":"","legend":"\u003cp\u003ePanels (A) display the pure colonies and (B) present their corresponding microscopic images at 100× magnification of strains CIP 82.92\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7316646/v1/38e4ccb02f46f205a3cc8f0b.png"},{"id":88812728,"identity":"dc7e9f88-3736-4eb1-b3ca-802c0d91da3f","added_by":"auto","created_at":"2025-08-11 15:51:45","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":154898,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree based on 16S rRNA gene sequences of strains CIP 82.92, constructed using the neighbor-joining method in MEGA X. Bootstrap values (1000 replicates) are indicated at branch nodes. The strain highlighted in green represents the reference strain CIP 82.92.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-7316646/v1/b9c5e0f03ce094e983ec4982.png"},{"id":88812730,"identity":"57deb038-c3c0-436f-af26-a39bdcdf54df","added_by":"auto","created_at":"2025-08-11 15:51:45","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":25485,"visible":true,"origin":"","legend":"\u003cp\u003eChanges in nitrogenase activity (nmol C₂H₄ h\u003csup\u003e-1 \u003c/sup\u003emL\u003csup\u003e-1\u003c/sup\u003e) and nitrogen concentration (mg L\u003csup\u003e-1\u003c/sup\u003e) of the selected strain at different inoculation times.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7316646/v1/d0f7ffa52c27e1c252cfe84c.png"},{"id":88814244,"identity":"d46f0ae9-0d1f-494d-b14e-8ad2e5fc2aca","added_by":"auto","created_at":"2025-08-11 16:09:00","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2107292,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7316646/v1/1f1de6c3-d961-4b4a-83ae-cd612e52dd1d.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Synergistic effect of Leclercia adecarboxylata CIP 82.92 and rice husk biochar on yield enhancement and cadmium reduction in mung bean","fulltext":[{"header":"Introduction","content":"\u003cp\u003eMung bean (\u003cem\u003eVigna radiata\u003c/em\u003e L.) is one of the most important pulse crops in many Asian countries, including Vietnam, India, China, and Thailand, owing to its high nutritional value, short growth cycle, nitrogen-fixing ability, and adaptability to various agro-ecological conditions (Hou et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). As a valuable source of plant-based protein, vitamins, minerals, and bioactive compounds, mung bean plays a crucial role in ensuring food security, improving soil fertility, and supporting smallholder farmers' livelihoods in resource-limited regions (Kessy et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In addition to its agronomic benefits, the crop contributes to sustainable agriculture by restoring soil nitrogen levels and reducing the need for synthetic fertilizers in crop rotation systems (Al-Shammary et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, in recent decades, the productivity and sustainability of mung bean cultivation have been increasingly threatened by a multitude of environmental stressors, most notably, climate change, soil degradation, and heavy metal contamination (Liu et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The intensification of agricultural production has led to the overuse of chemical fertilizers and pesticides, contributing to declining soil health, reduced microbial diversity, and increased environmental pollution (Xing et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Among these, cadmium (Cd) contamination has emerged as a particularly serious concern due to its high toxicity, persistence in soil, and ability to accumulate in edible plant parts (Chuong et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Cd contamination in agricultural soils, originating from industrial discharge, phosphate fertilizers, and irrigation with polluted water, poses a dual threat: it not only reduces crop productivity but also endangers human health through dietary intake (Ni\u0026ntilde;o-Savala et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Liang et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eMB, like many other legumes, is susceptible to Cd uptake and accumulation, which severely compromises seed quality and market value (Rehman et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Davidova et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Moreover, Cd stress negatively affects physiological and biochemical processes in plants, including nutrient uptake, photosynthesis, and enzymatic activity, ultimately leading to stunted growth and lower yields (Zhao et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rehman et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This situation underscores the urgent need for environmentally friendly and effective approaches to mitigate Cd toxicity in mung bean cultivation, particularly in regions where soil contamination is widespread and traditional remediation practices are either too costly or impractical (Umer Chattha et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zulfiqar et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In response to these challenges, the use of plant growth-promoting endophytic bacteria (PGPEB) in combination with organic soil amendments, such as biochar, has gained increasing attention as a sustainable strategy to enhance crop resilience, improve soil health, and reduce heavy metal uptake in plants (Kamyab et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Chuong et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Endophytic bacteria colonize plant tissues without causing disease and are known to confer multiple benefits to their host plants, including biological nitrogen fixation, phytohormone production, nutrient solubilization, and metal detoxification (Qadir et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Among various PGPEB, nitrogen-fixing strains are particularly valuable in legumes, as they reduce dependence on synthetic nitrogen inputs and enhance plant growth under stressful conditions (Chuong \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003ea).\u003c/p\u003e\u003cp\u003eRHB, a carbon-rich material produced by pyrolysis of biomass under limited oxygen conditions, has also been widely studied for its ability to immobilize heavy metals, improve soil structure, and support beneficial microbial communities. RHB is an abundant and low-cost by-product in many rice-producing countries (Asadi et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Its porous structure and high cation exchange capacity make it an effective soil amendment to reduce metal bioavailability and promote plant growth. When applied together, biochar and beneficial bacteria can create a synergistic effect, enhancing both microbial survival and plant performance under stress conditions (Kamyab et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Despite these promising findings, relatively few studies have examined the combined effect of indigenous endophytic bacteria and biochar on mung bean productivity and Cd mitigation under field conditions. Furthermore, the potential of \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e an emerging plant-associated bacterium with nitrogen-fixing potential remains largely unexplored in legume crops. \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e has been identified in various plant rhizospheres and endophytic environments, showing promise in promoting growth and stress tolerance in host plants. However, its application in Cd-contaminated soils, particularly in mung bean cultivation, is novel and warrants detailed investigation (Meng et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2025\u003c/span\u003e)\u003c/p\u003e\u003cp\u003eTo address these knowledge gaps, the present study was undertaken to isolate and identify indigenous endophytic bacteria from mung bean roots and evaluate their potential in combination with RHB to enhance plant growth, yield, and nutritional quality while reducing Cd accumulation under field conditions. Specifically, we hypothesized that (1) inoculation with strain \u003cb\u003eCIP 82.92\u003c/b\u003e would improve mung bean performance through biological nitrogen fixation and growth stimulation; (2) application of rice husk biochar would reduce Cd bioavailability and enhance soil fertility; and (3) the combined use of biochar and bacterial inoculation would exert a synergistic effect, leading to significant improvements in yield and reduction of Cd accumulation in mung bean tissues. This study provides new insights into the integrated use of indigenous microbial resources and organic amendments to rehabilitate Cd-contaminated agricultural soils and promote sustainable mung bean production. The results contribute to a growing body of evidence supporting the ecological benefits of nature-based solutions in mitigating heavy metal stress and reducing dependency on synthetic agrochemicals.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eCollection and treatment of MB root samples\u003c/h2\u003e\u003cp\u003eThe MB root samples were collected from MB cultivation fields in An Phu commune, An Giang province, Vietnam. This area is known to have Cd-contaminated soil, and previous studies have detected Cd accumulation in the stems, leaves, and seeds of the plants (Chuong et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The roots were carefully uprooted, placed in sterile plastic bags, and transported to the laboratory. Prior to isolation, the MB roots were surface sterilized by thoroughly washing them with tap water to remove soil and debris. The MB roots were then immersed in 70% ethanol for 3 minutes for partial sterilization, followed by treatment with 2.5% sodium hypochlorite for 5 minutes to eliminate most surface bacteria. Finally, the roots were rinsed again with 70% ethanol and several times with sterile distilled water to remove any remaining sterilizing agents. To verify the effectiveness of surface sterilization, sterilized root segments were placed on agar plates. The absence of bacterial growth after incubation indicated successful sterilization (GohilK PatelH et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saldierna Guzm\u0026aacute;n et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cb\u003eIsolation and Identification of\u003c/b\u003e Strain \u003cb\u003eCIP 82.92\u003c/b\u003e\u003c/p\u003e\u003cp\u003eFor the isolation of strain CIP 82.92, the surface-sterilized roots were homogenized and serially diluted up to 10⁻⁸. Aliquots (0.1 mL) from each dilution were spread onto yeast mannitol agar (YMA) plates and incubated at 32\u0026deg;C for 60 hours. Among the resulting colonies, ten isolates that exhibited morphological characteristics consistent with CIP 82.92 strain were selected for further study. These colonies were 3\u0026ndash;4 mm in diameter, had smooth and regular margins, and were large, rod-shaped, endospore-forming, and Gram-positive as confirmed by Gram staining and microscopic observation at 100X magnification (Nxumalo et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eMolecular identification of the ten selected isolates was conducted using 16S rRNA gene sequencing. Amplification was performed using the universal primers 27F (5\u0026prime;-AGAGTTTGATCCTGGCTCAG-3\u0026prime;) and 1492R (5\u0026prime;-GGTTACCTTGTTACGACTT-3\u0026prime;) (Dos Santos et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Hongmiao et al. 2021). The obtained sequences were compared with reference sequences in the GenBank database via BLAST. Phylogenetic analysis was carried out using MEGA 11 software, following the method of Saitou and Nei (\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1987\u003c/span\u003e) and Tamura et al. (\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), with 1,000 bootstrap replicates (Felsenstein, 1985). Ambiguous positions were excluded using the pairwise deletion option (Kumar et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). One colony was confirmed to be strain CIP 82.92, exhibiting 100% identity with reference strains. These sequences were subsequently submitted to the NCBI GenBank database and used to construct the phylogenetic tree (Fig.\u0026nbsp;1).\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eThermal adaptation\u003c/h3\u003e\n\u003cp\u003eThermal adaptation of strain CIP 82.92 was conducted in two phases. Initially, four YMA nutrient medium test tubes were prepared for each strain. Subsequently, strain CIP 82.92 streaked onto YMA agar in the tubes and incubated at four different temperatures: 30, 35, 40, and 45\u0026deg;C. Each thermal level was tested in four replicates, and colony development was observed over the course of one week (Van Chuong and Le Kim Tri \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eSalt adaptation\u003c/h3\u003e\n\u003cp\u003eThe salt tolerance of the selected strain was evaluated using the following method: First, four test tubes containing YMA agar medium were prepared for each salt concentration. Then, NaCl was added to achieve final concentrations of 1.0%, 2%, 3%, 4%, and 5%. Selected colonies of the strain CIP 82.92 were subsequently inoculated into each tube and incubated at 28\u0026deg;C, with four replicates per concentration. Observations were recorded after one week (Mustafa A and \u0026Ouml;zden E \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Van Chuong and Le Kim Tri \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eAmmonia production\u003c/h3\u003e\n\u003cp\u003eThe ability of the identified strain CIP 82.92 to produce ammonia was evaluated qualitatively. Individual colonies were inoculated into peptone water and incubated on YMA medium at 30\u0026deg;C for 60 to 80 hours. Ammonia production was confirmed by a color shift from brown to yellow upon the addition of Nessler\u0026rsquo;s reagent (Borah et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eNitrogenase activity\u003c/h3\u003e\n\u003cp\u003eNitrogenase activity was determined using the acetylene reduction assay following the method described by Puri et al. (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The strain CIP 82.92 was initially cultured in YMA liquid medium and incubated for 24 hours. The resulting culture was then used to inoculate a nitrogen-free liquid medium. A control was maintained using the nitrogen-free medium without the addition of the strain CIP 82.92. The cell density of the CIP 82.92 suspension, measured spectrophotometrically at 600 nm, reached 0.8 under incubation conditions of 30\u0026deg;C and 160 rpm (Soper et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eNitrogen concentration\u003c/h2\u003e\u003cp\u003eThe strain CIP 82.92 was grown in a nitrogen-free medium supplemented with 0.05% malate, serving as the main carbon source, and incubated at a constant temperature of 30\u0026deg;C. After the incubation period, the cultures were centrifuged at 3000 rpm for one minute to separate the biomass. The resulting supernatant was carefully collected and used for nitrogen content analysis. This determination was conducted according to the procedure described by Daoliang et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which allows for the accurate quantification of nitrogen fixed or released by the bacterial strains under nitrogen-limited conditions.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eDensity augmentation of strain CIP 82.92\u003c/h3\u003e\n\u003cp\u003eThe selected strains were cultured in sterile YMA medium and incubated at 32\u0026deg;C for 60 hours. After incubation, the cultures were centrifuged at 6,000 rpm for 5 minutes, and the supernatant was removed. The bacterial cells were washed with sterile saline to adjust the concentration to 10⁸ CFU mL⁻\u0026sup1;. Suspensions of the five selected strains were then used to prepare a 1.0% (v/v) inoculum for evaluating plant growth-promoting factors. In the field experiment, 100 mL of this suspension was applied to MB seeds prior to sowing (Song et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003ePreparation of MB variety\u003c/h3\u003e\n\u003cp\u003eThe DX 208 MB variety has a growth duration of 70 to 80 days, with a determinate growth habit. The average yield depends on the level of cultivation intensity. The seeds have a green, moldy appearance that meets consumer preferences. DX 208 exhibits synchronous maturity, making it convenient for harvesting, which is typically done 2 to 3 times per cropping season. The variety shows good lodging resistance and moderate to high resistance to Cercospora leaf spot and powdery mildew. DX 208 is well-suited for cultivation in delta and coastal regions within crop rotation systems (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://sonnptnt.ninhthuan.gov.vn\u003c/span\u003e\u003cspan address=\"https://sonnptnt.ninhthuan.gov.vn\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e)\u003c/p\u003e\u003cp\u003e\u003cb\u003eField Eexperimental design.\u003c/b\u003e\u003c/p\u003e\u003cp\u003eNPK fertilization for the 2023\u0026ndash;2024 and 2024\u0026ndash;2025 seasons: Fertilizers were applied at rates of 40 kg N, 60 kg P₂O₅, and 50 kg K₂O per hectare, divided into three stages as follows: (i) Basal application: The entire amount of P₂O₅, along with one-third of the N and one-third of K₂O, was applied before sowing; (ii) First topdressing (at the 3-true-leaf stage): An additional one-third of N and one-third of K₂O was applied; (iii) Second topdressing (25\u0026ndash;30 DAS): The remaining amounts of N and K₂O were applied (Trang and Chuong, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eField experiments were conducted in An Phu commune, An Giang province, Vietnam, located at approximately 10.845\u0026deg;N latitude and 105.080\u0026deg;E longitude, from October to January of the 2023\u0026ndash;2024 and 2024\u0026ndash;2025 seasons. The experimental site\u0026rsquo;s climate transitions from the rainy season to the dry season during this period. From October to November per year, the area experiences high rainfall and humidity, whereas December to January is dry with little rainfall and temperatures ranging from 22\u0026ndash;30\u0026deg;C, characterized by cool and stable weather. A randomized complete block design with two factors was used for both crop years (October to January of 2023\u0026ndash;2024 and 2024\u0026ndash;2025): Factor (1) with and without inoculation of strain CIP 82.92, and Factor (2) three levels of RHB application: 0.0, 5.0, and 10.0 t ha⁻\u0026sup1;. This resulted in six treatment combinations, each replicated four times. Inorganic fertilizers (40 kg N, 60 kg P₂O₅, and 50 kg K₂O per hectare) were uniformly applied to all treatments (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eTreatments of strain CIP 82.92 inoculation and different RHB rates in 2023\u0026ndash;2024 and 2024\u0026ndash;2025\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eTreatments\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eStrain CIP 82.92\u003c/p\u003e\u003cp\u003e(10\u003csup\u003e8\u003c/sup\u003e CFU mL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eRHB\u003c/p\u003e\u003cp\u003e(t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eChemical fertilizer\u003c/p\u003e\u003cp\u003e(kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e\u003cp\u003eYears\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"5\" rowspan=\"6\"\u003e\u003cp\u003e40N- 60P\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e5\u003c/sub\u003e-50 K\u003csub\u003e2\u003c/sub\u003eO\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMB6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThree pre-germinated mung bean seeds were sown per hole. Irrigation was carried out using water sourced from the Mekong River Delta. The experiment consisted of 6 treatments (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), each replicated four times, with a total experimental area of 480 m\u0026sup2; (2 m width \u0026times; 10 m length \u0026times; 4 replications \u0026times; 6 treatments). MB seeds were sown at a spacing of 25 cm \u0026times; 30 cm, with two seeds per hole, and thinned to retain one healthy plant 15 days after sowing (DAS). All treatments received chemical fertilizers at rates of 40 kg urea ha⁻\u0026sup1;, 60 kg P₂O₅ ha⁻\u0026sup1;, and 50 kg K₂O ha⁻\u0026sup1;, supplied by Binh Dien Fertilizer Company (Vietnam). Twenty-four soil samples were collected from the experimental treatments 15 days before the start of the experiment and analyzed, with the results presented in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. Agronomic parameters such as plant height, number of shoots, and chlorophyll content were monitored 65 days after sowing. Yield components, total yield, lipid and protein contents, as well as Cd accumulation in the stems, leaves, and seeds of MB were assessed annually at harvest. Cadmium concentrations were determined using atomic absorption spectrophotometry. Plant and soil analyses were conducted at the laboratories of An Giang University and Eurofins Company in Can Tho Province, Vietnam.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003ePhysicochemical properties of farmland soil and RHB prior to the experiment (n\u0026thinsp;=\u0026thinsp;24)\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eProperty\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eResult\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eProperty\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eResult\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSand (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e60.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTotal N (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.102\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSilt (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e36.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAvailable P (mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e72.7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eClay (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eExchangeable K (mg kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e106\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003epH soil\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSOM (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.21\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCadmium in soil (\u0026micro;g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e123\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCEC (cmol kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.02\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e2.11 Data collection and analysis\u003c/b\u003e\u003c/p\u003e\u003cp\u003eAgronomic and yield parameters in both years, including total biomass, number of filled pods, filled pod weight, 1000-seed weight, were recorded at harvest. Fresh pod yield (t ha⁻\u0026sup1;) was determined for each treatment in both experimental years. In addition, seed samples were subjected to biochemical analyses to quantify lipid, protein, and Cd concentration of MB stems, leaves and seeds, following standard analytical protocols. All collected data were statistically analyzed using Statgraphics XV and Microsoft Excel. Analysis of variance (ANOVA) was performed, and treatment means were compared using the Least Significant Difference (LSD) test at a 5% significance level (p\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eBiochemical and genetic profiling of strain CIP 82.92\u003c/h2\u003e\u003cp\u003eThe colony of strain CIP 82.92 was selected from 20 initially isolated pure colonies based on Gram staining results and morphological characteristics, followed by molecular identification. The selected strain, \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e CIP 82.92, was confirmed to share 100% genetic similarity with \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e NBRC 102595 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In the next step, its morphology was further examined to assess its nitrogen-fixing potential prior to conducting field experiments. The VITEK 2 system (BioM\u0026eacute;rieux, France) provided a 92% probability of identification and was further confirmed by Illumina-based genetic analysis. This system was employed to characterize the morphological and biochemical features of the four selected strains. Morphologically, prior to molecular identification, strain CIP 82.92 appeared as creamy white colonies with a diameter of approximately 3 mm, lacking pigmentation, and exhibiting shapes ranging from circular to irregular with smooth to undulate margins on YMA medium (Fig.\u0026nbsp;1). As shown in Fig.\u0026nbsp;1 shows that strain CIP 82.92 was identified as a Gram-positive, rod-shaped, motile bacterium, catalase-positive, and capable of aerobic growth at temperatures ranging from 20 to 50\u0026deg;C. Optimal growth occurred at pH 4\u0026ndash;8, within the same temperature range, and in media containing up to 5% NaCl. Strain B was primarily identified based on phenotypic characterization and partial results obtained from the VITEK 2 system, which are summarized in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. Furthermore, phylogenetic analysis based on 16S rRNA gene sequences, with a total branch length of 0.001, confirmed 100% genetic similarity with the reference sequence of \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e NBRC 102595, as illustrated in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThe physical and chemical characterization of strains CIP 82.92 was determined using the VITEK 2 analyzer\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBiochemical test\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003estrains CIP 82.92\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBiochemical test\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003estrains CIP 82.92\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBeta\u0026ndash;Xyloidine\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNaCl (1\u0026ndash;5%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e++\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eD\u0026ndash;Mannose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTemperature (20-50\u003csup\u003e0\u003c/sup\u003eC)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eD\u0026ndash;Glucose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003epH (4.0\u0026ndash;8.0)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e++\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eD\u0026ndash;Galactose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCitrate use\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eD\u0026ndash;Ribose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eβ \u0026ndash; Glucosidase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCatalase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e++\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMannitol\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOxidase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eRaffinose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStarch hydrolysis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eGlucose oxidation\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003e(-): negative reaction; (+) weak reaction; (++) strong reaction\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003e3.2 Assessment of ammonia activity and concentration\u003c/b\u003e\u003c/p\u003e\u003cp\u003eThe acetylene reduction to ethylene (C₂H₄) process, monitored via the C₂H₂ gas injection system, initially exhibited a short lag phase after acetylene introduction and gradually stabilized over the 72-hour incubation period. As shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e3\u003c/span\u003e, both nitrogenase activity and nitrogen concentration increased progressively with inoculation time. Specifically, the nitrogenase activity began at a low level (approximately 20 nmol C₂H₄/h/mL at 8 hours) and steadily rose, peaking at around 380 nmol nmol C₂H₄ h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e mL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003eafter 72 hours. Similarly, nitrogen concentration measured by the Kjeldahl method followed a comparable upward trend, increasing from below 50 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e at 8 hours to over 420 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e at 72 hours. Findings confirm that the selected strain CIP 82.92 actively participates in biological nitrogen fixation, with clear evidence of sustained nitrogenase enzyme activity over time. The high correlation between ethylene production and nitrogen accumulation suggests that this strain maintains robust diazotrophic capacity under laboratory conditions.\u003c/p\u003e\u003cp\u003e\u003cb\u003eEffect of strain CIP 82.92 and RHB on MB agronomic traits\u003c/b\u003e.\u003c/p\u003e\u003cp\u003eThe data presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e clearly demonstrates the positive influence of both strain CIP 82.92 and RHB on the agronomic traits of mung bean, including plant height, number of branches per plant, and chlorophyll index, measured at 65 DAS. Across both years, inoculation with strain CIP 82.92 significantly improved all measured parameters. In the 2024\u0026ndash;2025 season, inoculated plants reached a mean height of 67.5 cm compared to 60.5 cm in non-inoculated controls, while branch number increased from 12.0 to 17.9, and chlorophyll index rose dramatically from 43.9 to 57.6.\u003c/p\u003e\u003cp\u003eApplication of RHB at increasing rates also significantly improved mung bean growth. The highest application rate (10.0 t ha⁻\u0026sup1;) resulted in the greatest increases in all parameters in both years, especially in 2024\u0026ndash;2025 where plant height and chlorophyll index peaked at 74.5 cm and 59.6, respectively. Significant interaction effects [F(A\u0026times;B)] were observed, indicating a synergistic relationship between CIP 82.92 and RHB. For example, combined treatment consistently resulted in greater improvements than either factor alone. Notably, the improvements were more pronounced in the 2024\u0026ndash;2025 season across all treatments. Biomass accumulation is a direct indicator of plant vigor and photosynthetic efficiency. According to the results, both the application of Bacillus strain CIP 82.92 and rice husk biochar (RHB) significantly increased the biomass of mung bean in both cropping years, with a notably stronger response in the 2024\u0026ndash;2025 season.\u003c/p\u003e\u003cp\u003eInoculation with CIP 82.92 increased biomass from 20.2 g plant⁻\u0026sup1; to 25.5 g plant⁻\u0026sup1; in 2023\u0026ndash;2024, and from 20.1 g to 30.5 g plant⁻\u0026sup1; in 2024\u0026ndash;2025. The more substantial increase in the second year (a 52% rise versus 26% in the first year) suggests that environmental conditions or cumulative soil microbiome enhancement may have amplified the bacterial effect. RHB at 10.0 t ha⁻\u0026sup1; led to the highest biomass in both seasons, increasing from 28.7 g (2023\u0026ndash;2024) to 40.8 g plant⁻\u0026sup1; (2024\u0026ndash;2025). Even at 5.0 t ha⁻\u0026sup1;, RHB significantly enhanced plant biomass over the control. The combined effect of CIP 82.92 and RHB was statistically significant [F(A\u0026times;B)] at p\u0026thinsp;\u0026le;\u0026thinsp;0.01, indicating a synergistic interaction. Notably, the biomass response was consistently greater in 2024\u0026ndash;2025 across all treatments, suggesting long-term soil conditioning or climatic factors improved microbial establishment and root-soil interactions.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eEffect of strains CIP 82.92 and RHB on MB-agronomic components at 65 DAS\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"12\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eFactors\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003ePlant height (cm)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eBranch number (branches plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"5\" nameend=\"c10\" namest=\"c6\"\u003e\u003cp\u003eChlorophyll index\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eBiomass\u003c/p\u003e\u003cp\u003e(g plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"11\" nameend=\"c12\" namest=\"c2\"\u003e\u003cp\u003eYears\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"10\" nameend=\"c10\" namest=\"c1\"\u003e\u003cp\u003eStrains CIP 82.92 (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e61.5b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e60.5b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e12.3b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12.0b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e44.9b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e43.9b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e20.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e20.1b\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e64.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e67.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e15.9a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.9a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e47.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e57.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e25.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e30.5a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"10\" nameend=\"c10\" namest=\"c1\"\u003e\u003cp\u003eRHB application (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e61.3c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e61.3b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e12.75b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12.7b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e44.0b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e42.0b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e20.1b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e19.6c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e63.1b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e73.1ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e15.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e48.3a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e58.3a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e28.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e33.5b\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e10.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e64.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e74.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e15.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.5a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e49.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e59.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e28.7a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e40.8a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A x B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"12\"\u003eNote: \u0026plusmn; indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at the 1% levels.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eEffect of strain CIP 82.92 and RHB on MB yield traits and productivity\u003c/h2\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eEffect of strains CIP 82.92 and RHB on MB yield trait and productivity\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"12\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eFactors\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003ePod number\u003c/p\u003e\u003cp\u003e(Pods per plant)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eFilled pod weight\u003c/p\u003e\u003cp\u003e(g plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"5\" nameend=\"c10\" namest=\"c6\"\u003e\u003cp\u003eWeight of 1, 000 seeds (g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eFresh yield\u003c/p\u003e\u003cp\u003e(t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"11\" nameend=\"c12\" namest=\"c2\"\u003e\u003cp\u003eYears\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"10\" nameend=\"c10\" namest=\"c1\"\u003e\u003cp\u003eStrains CIP 82.92 (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e35.7b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e30.1b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e18.5b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e16.5b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e35.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e32.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e1.5c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e1.61c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e45.7a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e54.1a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e25.8a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e27.8a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e39.1a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e41.1a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e2.1a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2.42a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"10\" nameend=\"c10\" namest=\"c1\"\u003e\u003cp\u003eRHB application (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e33.4c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e32.1c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e16.8c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.8b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e33.2c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e33.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e1.3c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e1.9c\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e43.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e50.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e24.7b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e28.7a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e45.2b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e49.2a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e2.0a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2.21b\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e10.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e45.7a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e55.6a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e27.0a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e28.0a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e47.4a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e49.4a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e1.97a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e2.54a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A x B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"12\"\u003eNote: \u0026plusmn; indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at P\u0026thinsp;\u0026gt;\u0026thinsp;0.01.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e proves that application of CIP 82.92 significantly increased pod numbers from 35.7 to 45.7 pods/plant in 2023\u0026ndash;2024 and even more so in 2024\u0026ndash;2025, from 30.1 to 54.1 pods/plant, reflecting a 79.7% increase in the second year. Similarly, RHB at 10.0 t ha⁻\u0026sup1; resulted in the highest pod numbers (45.7 and 55.6 pods plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e in 2023\u0026ndash;2024 and 2024\u0026ndash;2025, respectively). Significant enhancements were recorded with both treatments. Strain CIP 82.92 increased filled pod weight by 39% (2023\u0026ndash;2024) and 68% (2024\u0026ndash;2025). RHB also showed consistent increases, especially at 5.0\u0026ndash;10.0 t ha⁻\u0026sup1; (up to 28.7 g plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). The MB pod number and weight are critical yield components and market determinants. CIP 82.92 increased 1000-seed weight by about 11% (2023\u0026ndash;2024) and 28% (2024\u0026ndash;2025).\u003c/p\u003e\u003cp\u003eFresh yield is the ultimate productivity measure. Inoculation with CIP 82.92 raised yield from 1.5 to 2.1 t ha⁻\u0026sup1; in the first year and from 1.61 to 2.42 t ha⁻\u0026sup1; in the second. RHB significantly enhanced yield, especially at 10.0 t ha⁻\u0026sup1;, reaching 2.54 t ha⁻\u0026sup1;, the highest recorded.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eImpact of strain CIP 82.92 and RHB nutrient compositions and Cd accumulation in MB seeds\u003c/h2\u003e\u003cp\u003eBoth strain CIP 82.92 and RHB significantly increased the lipid and protein contents of MB seeds over the two years. Notably, CIP 82.92 increased lipid content from 25.2\u0026ndash;26.8% in 2023\u0026ndash;2024 and 25.3\u0026ndash;27.8% in 2024\u0026ndash;2025, while protein content improved from 17.3\u0026ndash;18.0% and 17.1\u0026ndash;19.0%, respectively. RHB application showed a dose-dependent improvement, especially at 10 t ha⁻\u0026sup1;, with lipid content reaching 31.9% and 32.3%, and protein content 18.2% and 19.2% in the two years, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eCd accumulation in both plant parts was significantly reduced by treatments with CIP 82.92 inoculation and RHB application. inoculation of strain CIP 82.92 reduced Cd in seeds from 61.7 to 36.1 \u0026micro;g kg⁻\u0026sup1; (2023\u0026ndash;2024) and from 77.6 to 23.4 \u0026micro;g kg⁻\u0026sup1; (2024\u0026ndash;2025). A 62\u0026ndash;70% reduction, confirming its Cd-immobilizing and phytoprotective properties. The Cd concentrations in stems and leaves also dropped significantly under CIP 82.92, indicating systemic mitigation. RHB application at 5.0 t ha⁻\u0026sup1; reduced seed Cd to 23.4 and 11.9 \u0026micro;g kg⁻\u0026sup1;, while 10.0 t ha⁻\u0026sup1; lowered it to 37.6 \u0026micro;g kg⁻\u0026sup1; in 2023\u0026ndash;2024 and undetectable levels in 2024\u0026ndash;2025. Importantly, the interaction effect [F(A\u0026times;B)] was significant for Cd reduction, particularly in seeds, indicating that co-application of Strains CIP 82.92 and RHB is more effective than either alone. The co-application of CIP 82.92 and RHB significantly enhanced the nutritional quality of mung bean seeds while drastically reducing cadmium accumulation, particularly in the second year. These findings support the integration of endophytic bacteria and biochar as a sustainable, eco-friendly strategy for safe legume production on Cd-contaminated soils (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eEffect of strain CIP 82.92 and RHB on nutrient traits and Cd accumulation in MB seeds\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"11\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eFactors\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eLipid\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eProtein\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"6\" nameend=\"c11\" namest=\"c6\"\u003e\u003cp\u003eCd concentration (\u0026micro;g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e\u003cp\u003eSeeds (%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c9\" namest=\"c6\"\u003e\u003cp\u003eStems and leaves\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c11\" namest=\"c10\"\u003e\u003cp\u003eSeeds\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"10\" nameend=\"c11\" namest=\"c2\"\u003e\u003cp\u003eYears\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e2023\u0026ndash;2024\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e2024\u0026ndash;2025\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"9\" nameend=\"c9\" namest=\"c1\"\u003e\u003cp\u003eStrains CIP 82.92 (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e25.2 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e25.3b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e17.3 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.1 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e90.9 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e108a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e61.7 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e77.6 a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eYes\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e26.8a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e27.8a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e18.0 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e19.0 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e62.2 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e53.0b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e36.1b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e23.4 b\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"9\" nameend=\"c9\" namest=\"c1\"\u003e\u003cp\u003eRHB application (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e0.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e28.4c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e27.4b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e16.9b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e17.3b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e159 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e117 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e77.6 a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e87.6a\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e31.1b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e32.1a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e17.9a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e19.9a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e65.2 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e63.0b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e23.4 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e11.9b\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e10.0 t ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e31.9a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e32.3a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e18.2a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e19.2a\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e64.7 b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e59.1 c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e37.6 c\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003eundetected\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eF (A x B)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e\u003cp\u003ens\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u003cp\u003e**\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"11\"\u003eNote: \u0026plusmn; indicates the standard deviation. Values sharing the same letter within a column do not differ significantly (ns), while ** denote significant differences at P\u0026thinsp;\u0026lt;\u0026thinsp;0.01.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e\u003ch2\u003eBiochemical and genetic profiling of strain CIP 82.92\u003c/h2\u003e\u003cp\u003eThe strain CIP 82.92, a Gram-positive, rod-shaped bacterium, was isolated from MB roots cultivated in cadmium-contaminated soils using YMA medium. Biochemical assays confirmed positive catalase and oxidase activities, consistent with the typical characteristics of endophytic nitrogen-fixing bacteria reported in recent studies (Hossain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The strain demonstrated moderate salt tolerance, thriving in media containing 1\u0026ndash;5% NaCl, which aligns with findings that most endophytes adapt well to low-to-moderate salinity conditions (Chandran et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Khushboo et al. 2023; Falf\u0026aacute;n-Cort\u0026eacute;s et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Molecular identification based on 16S rRNA gene sequencing revealed 100% similarity with known nitrogen-fixing bacteria, confirming the taxonomic position of the strain as \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e, a species recognized for its growth-promoting capacity and cadmium resistance (Wang et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The optimal growth temperature ranged from 20\u0026deg;C to 50\u0026deg;C, with 37\u0026deg;C being ideal for symbiotic nitrogen fixation, consistent with earlier reports on endophyte thermotolerance (Van Chuong and Le Kim Tri \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Nguyen Van \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e\u003ch2\u003eEffect of strain CIP 82.92 and RHB on MB agronomic traits\u003c/h2\u003e\u003cp\u003eRecent studies have emphasized that \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e not only plays a crucial role in nitrogen fixation under abiotic stress conditions but also promotes plant development via phytohormone production, phosphate solubilization, and heavy metal resistance demonstrating its potential as a biofertilizer in sustainable agriculture (Woo et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These results are in accordance with previous findings confirming the effectiveness of the acetylene reduction assay (ARA) as a reliable indicator of nitrogenase activity and biological nitrogen fixation capacity (Aasfar et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This supports prior findings that biochar enhances soil structure, cation exchange capacity, and microbial habitat (Torita et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This synergy suggests that biochar not only improves the physical-chemical properties of soil but also serves as a carrier or habitat for beneficial microbes, enhancing their survival and colonization efficiency. Similar interactions have been reported by Cao et al. (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), who observed that biochar-amended soils facilitated root colonization by endophytic bacteria, leading to improved nutrient uptake and growth. This aligns with the nitrogen-fixing potential of CIP 82.92, as previously confirmed through acetylene reduction assays (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e3\u003c/span\u003e), and suggests more efficient nitrogen use efficiency when both amendments are applied in combination (Barbosa et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The combined application of strain CIP 82.92 and RHB significantly enhanced MB growth attributes in both years, with stronger responses observed in the second season. These results highlight the potential of integrating PGPR and biochar as a sustainable and synergistic strategy for legume crop improvement.\u003c/p\u003e\u003cp\u003eThe high nitrogenase activity of strain CIP 82.92, as indicated by the ARA, reflects its strong potential for biological nitrogen fixation. This observation is consistent with recent reports on \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e being isolated from leguminous crops (Meng et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Woo et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The elevated nitrogen levels recorded in the essay further confirm the strain\u0026rsquo;s effectiveness in enhancing nitrogen bioavailability, is a key factor in crop productivity. This effect likely stems from the strain's ability to adapt to the rhizosphere environment and to establish symbiotic interactions with legume roots (Concha et al. 2020; Chuong et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Its nitrogen-fixing efficiency is comparable to, or even surpasses, other known endophytic diazotrophs, underscoring its potential as a biofertilizer candidate (Figiel et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e\u003ch2\u003eEffect of strain CIP 82.92 and RHB on MB yield trait and yield\u003c/h2\u003e\u003cp\u003eThe application of strain CIP 82.92 in combination with RHB significantly improved nutrient availability and vegetative growth in MB. This is supported by recent studies highlighting RHB\u0026rsquo;s role in enhancing soil structure and nutrient uptake in legumes (Ben Gaied et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The observed plant growth-promoting effects of CIP 82.92 may be attributed to mechanisms such as phytohormone production, phosphate solubilization, and nitrogen fixation, as also reported by Etesami (\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and Flores et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The interactive effect between the microbial inoculant and organic amendment underscores the synergistic benefits of integrating beneficial microbes with organic inputs, a strategy widely advocated in integrated nutrient management systems (Folina et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Liang et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eField trials conducted over two consecutive years revealed that combined application of CIP 82.92 and RHB at 10 t ha⁻\u0026sup1; significantly enhanced MB yield and quality parameters compared to uninoculated or singly treated controls. Filled pod number reached 176 pods per plant and filled pod weight increased to 196 g per plant, representing 31% and 65% increases, respectively, over untreated controls. In contrast, the number and weight of unfilled pods decreased, indicating more efficient reproductive development and resource partitioning (LSD\u0026thinsp;\u0026le;\u0026thinsp;0.01). These results are consistent with prior studies highlighting the role of endophytic bacteria and biochar in improving flower development, nutrient acquisition, and carbon allocation to reproductive organs (Villa-Parejo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eRHB applications were also shown to improve soil physical properties and nutrient retention, supporting better seed development (Trozzo et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Thousand-seed weight increased to 47.4\u0026ndash;49.4 g under the 10t RHB ha⁻\u0026sup1; treatment, reflecting improved assimilate allocation for seed filling (Zhu et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These findings are consistent with recent reports demonstrating that RBH enhances nutrient availability and soil microbial activity, leading to increased growth and yield of leguminous crops (Abdul-Aziz et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Biochar is known to enhance aeration, water retention, and microbial habitat quality in soil, particularly after multi-season application (Kumar et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eRecent studies further indicate that RHB combined with endophytic nitrogen-fixing bacteria significantly improves nodule formation and legume productivity, including in mung bean compared to treatments with low fertilizer or no inoculants (Linh et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Notably, microbial inoculation often exerts greater influence on yield improvement than nitrogen fertilizers alone, emphasizing the value of biological inputs in sustainable legume production systems (L\u0026oacute;pez-Rom\u0026aacute;n et al.2025). In addition to boosting yield, the RHB-bacteria combination enhances soil health by increasing organic matter and beneficial microbial populations, which support long-term soil fertility (Ali et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe integration of CIP 82.92 with RHB offers a promising strategy for increasing mung bean yield and seed protein content while concurrently reducing cadmium accumulation in edible plant parts. This approach contributes to sustainable agriculture through improved soil fertility, reduced environmental impact of synthetic fertilizers, and potential biocontrol of root diseases such as \u003cem\u003ePhytophthora\u003c/em\u003e root rot via microbial antagonism and enhanced plant health (Woo et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The use of biofertilizer and RHB combinations is recommended for MB cultivation on nutrient-depleted and Cd-contaminated soils to maximize productivity, farmer income, and environmental sustainability. The observed effects may result from the strain\u0026rsquo;s nitrogen-fixing and phosphate-solubilizing capabilities, which facilitate more efficient nutrient uptake consistent with recent findings on endophytic bacteria in sustainable farming systems (Dadhich et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec19\" class=\"Section2\"\u003e\u003ch2\u003eEffect of strain CIP 82.92 and RHB on MB nutrition and Cd accumulation\u003c/h2\u003e\u003cp\u003eThese results indicate that CIP 82.92 promotes plant growth, possibly through enhanced nitrogen fixation and phytohormone synthesis consistent with previous studies on beneficial endophytes such as \u003cem\u003eBacillus aryabhattai\u003c/em\u003e and \u003cem\u003eBacillus subtilis\u003c/em\u003e (Radhakrishnan et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Chuong et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003eb). Biochar\u0026rsquo;s known capacity to improve soil aeration, moisture retention, and nutrient availability also provides favorable conditions for microbial colonization (Chen et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The synergistic effect of biochar and PGPR has been shown to increase photosynthetic efficiency and biomass accumulation in legumes (Zhang et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Stronger plant responses observed in the second year (2024\u0026ndash;2025) may reflect cumulative benefits from biochar conditioning, microbial establishment, and improved soil health. These findings are supported by studies reporting that repeated applications of biochar and PGPR enhance rhizosphere stability and functional resilience (Zhang et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Trang and Chuong, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe greater reduction in Cd uptake and higher nutrient concentrations observed in 2024\u0026ndash;2025 may be due to long-term improvements in soil structure, microbial colonization, and Cd immobilization. This supports biochar\u0026rsquo;s role in enhancing nutrient availability and microbe\u0026ndash;root interactions, as previously reported (Ahmad et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Potential mechanisms include extracellular polymeric substance (EPS) production, biosorption, and modified cadmium translocation pathways\u0026mdash;consistent with recent studies on endophytic bacteria (Haider et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Collectively, these findings confirm that biochar functions as a strong sorbent, binding heavy metals in soil and reducing plant uptake (Ma et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eBased on the conducted field experiments over two consecutive seasons (2023\u0026ndash;2024 and 2024\u0026ndash;2025) in Cd-contaminated soils of An Phu commune. The synergistic application of strain CIP 82.92 and RHB significantly enhanced MB growth, yield, and seed quality while reducing Cd accumulation. The inoculation with strain CIP 82.92 improved plant height, branch number, chlorophyll content, biomass accumulation, pod numbers, filled pod weight, and 1,000-seed weight compared to uninoculated controls. Increasing RHB rates (up to 10 t ha⁻\u0026sup1;) also positively affected all growth and yield parameters, with maximal improvements observed under combined treatments, indicating a strong synergistic interaction. Importantly, the combined treatment substantially reduced cadmium concentrations in seeds and aerial plant parts, enhancing food safety. Seed lipid and protein contents were elevated with both CIP 82.92 and RHB treatments, further improving seed nutritional value. The beneficial effects are attributed to enhanced nitrogen fixation, phytohormone production from CIP 82.92, and improved soil physicochemical properties and microbial habitats provided by RHB. The more pronounced benefits observed during the second year suggest cumulative improvements in soil health and microbial establishment. This study demonstrates the potential of integrating endophytic bacteria and biochar amendments as an effective agronomic strategy to increase legume productivity and mitigate heavy metal stress in contaminated soils.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor contributions:\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eTran Thanh Liem, Phan Tran Hai Dang, Tran Le Kim Tri and Nguyen Ngoc Phuong Trang conducted experiments, collected data, performed chemical analyses, and assisted with statistical analysis. Nguyen Van Chuong conceived and designed the study, supervised the project, analyzed data, drafted and finalized the manuscript for submission.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eNo funding was received to assist with the preparation of this manuscript\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e No datasets were generated or analyzed during the current study\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability\u003c/strong\u003e Not applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclarations Ethical approval\u003c/strong\u003e Not applicable.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate\u003c/strong\u003e Not applicable.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e Not applicable.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e the authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAasfar A, Meftah Kadmiri I, Azaroual SE, Lemriss S, Mernissi NE, Bargaz A, Zeroual Y, Hilali A (2024) Agronomic advantage of bacterial biological nitrogen fixation on wheat plant growth under contrasting nitrogen and phosphorus regimes. 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Plants 12:3147. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi:10.3390/plants12173147\u003c/span\u003e\u003cspan address=\"https://doi:10.3390/plants12173147\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Biochar, Cadmium, Indigenous bacteria, Nitrogen-fixing, Strain CIP 82.92","lastPublishedDoi":"10.21203/rs.3.rs-7316646/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7316646/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMung bean (\u003cem\u003eVigna radiata\u003c/em\u003e L.) is a vital legume crop widely grown in Asia, valued for its nutritional qualities and nitrogen-fixing ability. However, its productivity is challenged by cadmium (Cd) contamination in soils, which poses risks to crop yield and food safety. This study aimed to isolate indigenous endophytic bacteria capable of promoting mung bean growth while reducing Cd uptake, and to evaluate the combined effect of bacterial inoculation with rice husk biochar (RHB) in field trials conducted over two consecutive seasons (2023\u0026ndash;2024 and 2024\u0026ndash;2025) in Cd-contaminated soils of An Giang province, Vietnam. \u003cem\u003eLeclercia adecarboxylata\u003c/em\u003e CIP 82.92 (strain CIP) was isolated from mung bean (MB) roots and characterized for morphological, biochemical, and molecular features, confirming its identity and nitrogen-fixing potential. Field experiments used a randomized complete block design with two factors: inoculation with CIP 82.92 and three RHB application rates (0, 5, and 10 t ha⁻\u0026sup1;), with chemical fertilizers uniformly applied. Results demonstrated that inoculation with CIP 82.92 significantly enhanced plant growth parameters, biomass, and yield components, while RHB application further improved these effects, showing a synergistic interaction. Importantly, Cd accumulation in seeds and aerial parts was markedly reduced by combined treatment, alongside improvements in seed protein and lipid content. The beneficial outcomes were more pronounced in the second year, indicating cumulative soil health benefits. This study highlights the integrated use of strain CIP 82.92 and RHB as an effective, eco-friendly approach to increase mung bean productivity and food safety in Cd-contaminated agricultural systems.\u003c/p\u003e","manuscriptTitle":"Synergistic effect of Leclercia adecarboxylata CIP 82.92 and rice husk biochar on yield enhancement and cadmium reduction in mung bean","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-08-11 15:43:40","doi":"10.21203/rs.3.rs-7316646/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"db9881ec-96af-4ce8-804e-3aba23cb0156","owner":[],"postedDate":"August 11th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-08-11T15:43:42+00:00","versionOfRecord":[],"versionCreatedAt":"2025-08-11 15:43:40","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7316646","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7316646","identity":"rs-7316646","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}